ˆ (a , ˆ ) ˘old.cepl.rssi.ru/bio/tan/109 morocco caves 2014.pdf · & tanasevitch (1996), for...

Linyphiid spiders (Araneae, Linyphiidae) from caves of Morocco

Andrei v. TANAseviTCHinstitute of ecology and evolution, Russian Academy of sciences, leninsky Prospect, 33, Moscow 119071, Russia. e-mail: [email protected]

Linyphiid spiders (Araneae, Linyphiidae) from caves of Morocco. -Nine species of linyphiid spiders are recorded from caves of Morocco, fourof which are described as new to science: Diplocephalus inanis sp. n.,Lepthyphantes s. lat. longipedis sp. n., Lepthyphantes s. lat. taza sp. n. andMegalepthyphantes brignolii sp. n.

Keywords: Arachnida - North Africa - taxonomy - new species - new records.

iNTRoduCTioNA small collection of linyphiid spiders from caves of Morocco is stored in the

Muséum d’histoire naturelle de genève. This material was studied by Paolo MarcelloBrignoli in the mid 80s of the last century, and the corresponding manuscript was almost finished. Regrettably, Brignoli passed away in 1986, and his manuscript withthe original descriptions cannot be found and is probably lost. only the line drawingsin ink, produced by the geneva Museum on the basis of Brignoli's pencil sketches, have remained and some of them are used in the current paper.

The collection contains one nominal species (Lepthyphantes s. lat. maurusius

Brignoli, 1978) and eleven species were labeled by Paolo Brignoli as new to science.Today, after 30 years, it is clear that four of these “new species” had already been described at that time: L. s. lat. aelleni denis in denis & dresco, 1957, L. s. lat. brevi-

hamatus Bosmans, 1985, L. s. lat. longihamatus Bosmans, 1985 and Tenuiphantes

tenuis (Blackwall, 1852), but four others are really new and described below. Threespecies from the collection represented by females only and i disregard them becausethe absence of corresponding males makes their identity unclear. This is a female belonging to Araeoncus simon, 1884 or to Diplocephalus Bertkau in Förster &Bertkau, 1883, and a few female specimens of two species, probably belonging to theafer species-group of Lepthyphantes s. lat. (see Brignoli, 1971; saaristo & Tanasevitch,1993).

All “Lepthyphantes species” mentioned in the current paper are cited as“Lepthyphantes s. lat.” and they do not belong to the genus Lepthyphantes Menge,1866 sensu saaristo & Tanasevitch (1996). Recently, the genus Lepthyphantes wasreassessed and limited to five species only (op. cit.). More than 400 species previouslyplaced in Lepthyphantes were transferred to other genera, but about 160 species aretemporarily left in Lepthyphantes until their taxonomic position is reassessed.

Revue suisse de Zoologie 121 (2): 277-290; juin 2014

Manuscript accepted 05.03.2014

MATeRiAl ANd MeTHodsThe present paper treats linyphiid spiders collected in Moroccan caves in 1974,

1978 and 1979 by P. strinati, B. Hauser and v. Aellen, and in 1982 by a team of speleologists from lyon (see gilbert, 1983). All material is kept at the Muséum d’histoire naturelle, geneva, switzerland (MHNg). sample numbers are given insquare brackets.

The majority of the figures used in this paper were made by Paolo Brignoli, idrew a few additional elements, made some corrections and provided figures with ab-breviations.

The terminology of genitalic structures in Micronetinae follows that of saaristo& Tanasevitch (1996), for erigoninae it mainly follows that of Hormiga (2000). Thechaetotaxy of erigoninae is given in a formula (e.g., 2.2.1.1) which refers to the number of dorsal spines on tibiae i-iv. For Micronetinae the chaetotaxy is given in adifferent formula, e.g., Ti i: 2-1-1-2(1), which means that tibia i has two dorsal spines,one prolateral spine, one retrolateral spine, and two or one ventral spines (the apicalspines are disregarded). The sequence of leg segment measurements is as follows: femur + patella + tibia + metatarsus + tarsus. All measurements are given in milli -metres. Brignoli’s figures were not drawn to scale, but all of my own illustrations aresupplied with scale bars.

The following abbreviations are used in the text and figures: BC - bursa copu-latrix; dPs - distal part of scape; dRA - distal radical apophysis; dsA - distal supra-tegular apophysis; e - embolus; eB - embolus base; eg - entrance groove; eP - em-bolus proper; Fe - femur; l - lamella characteristica; lW - lateral wall; lWP - lateralwall process; MHNg - Muséum d’histoire naturelle, geneva, switzerland; MPs -middle part of scape; Mt - metatarsus; PMP - posterior median plate; Ps - proscape;PsB - proscape base; R - radix; st - stretcher; TA - terminal apophysis; Th - thumb; Ti- tibia; Tmi - relative position of trichobothrium on metatarsus i.

ResulTs

Diplocephalus inanis sp. n. Figs 1-8HoloTyPe: �; MoRoCCo, Middle Atlas Mts, south west of Taza, Châra, near

AinTeslit, “ifri Tselet” Cave, 1250 m a.s.l.; 3.vi.1978; leg. P. strinati [Mar78/14].PARATyPe: 1 �; collected together with the holotype.

eTyMology: The specific epithet is a latin adjective; one of its many meaningsis “eyeless”.

diAgNosis: The new species is characterized by the absence of eyes in bothsexes, by the absence of any process on the male palpal tibia, as well as by the thickseminal ducts which are clearly visible through both parts of the epigynal ventral plate.

desCRiPTioN: Male (holotype), partly damaged. Total length 1.30. Carapace0.58 long, 0.48 wide, pale brown, almost yellow. Cephalic part slightly elevated, sulcipresent, narrow, almond-shaped; eyes totally reduced (Figs 1-3). Chelicerae 0.23 long,unmodified. legs mostly broken off. Fei 0.68, Feiv 0.80 long. Chaetotaxy unknown.Palp (Figs 5-7): Palpal tibia thickened, without apophyses but with a wide distal lobe.distal suprategular apophysis developed as a long stripe, narrowing in distal part.

A. v. TANAseviTCH278

embolic division large, distal radical apophysis with two pointed apophyses of different lengths. embolus relatively short, directed backwards, strongly curved distally. Abdomen about 0.83 long, 0.50 wide, pale, almost white.

Female. Total length ca 1.60. Carapace 0.65 long, 0.48 wide, pale brown, un-modified, eyes totally reduced (Fig. 4). Chelicerae 0.50 long. legs pale brown. leg i2.57 long (0.75+0.20+0.65+0.55+0.42), leg iv 2.59 long (0.78+0.18+0.68+0.67+0.38). Chaetotaxy 2.2.1.1. Metatarsus iv without trichobothrium. Tmi 0.36.Abdomen 0.95 long, 0.58 wide, pale, almost white. epigyne (Fig. 8): Both parts of bisected ventral plate wide apart proximally, virtually in contact with each other distally. dorsal plate shaped like a narrow rectangle. entrance ducts thick and well visible through both parts of ventral plate in median fissure area.

ReMARks: There is only a single known eyeless species of Diplocephalus

Bertkau in Förster & Bertkau, 1883, D. caecus denis, 1952, which was described froma male from a cave in Rochefort, Belgium. Diplocephalus inanis sp. n. can easily be

279liNyPHiid sPideRs FRoM MoRoCCo

Figs 1-4Diplocephalus inanis sp. n., � holotype (1-3) and � paratype (4). (1) Carapace, lateral view.(2) same, dorsal view. (3, 4) same, frontal view.

distinguished from that species by the absence of any process on the palpal tibia, aswell as by some other details of the palp.

The absence of any apophyses on the palpal tibia is a rare phenomenon amongmembers of Diplocephalus, and elsewhere only found in the North-African D. mysta-

cinus (simon, 1884), which occurs in Algeria and Tunisia (and which also has a distallobe), and in the european-Mediterranean D. graecus (Cambridge, 1872). The new


Figs 5-8Diplocephalus inanis sp. n., � holotype (5-7) and � paratype (8). (5-6) left palp, retrolateraland prolateral view, respectively. (7) Palpal tibia, dorsal view. (8) Cleared epigyne, ventral view.

species clearly differs from both species by the absence of eyes and by details of thegenitalia.

disTRiBuTioN: known from the type locality only.

Lepthyphantes s. lat. aelleni denis in denis & dresco, 1957 Figs 9-13MATeRiAl: 1 �; MoRoCCo, Middle Atlas Mts, Taza, near daya Chiker, “gouffre du

Friouato” Cave, 1450 m a.s.l.; 1.vi.1978; leg. P. strinati [Mar 78/6]. – 1 �; same data,1.vi.1978; leg. B. Hauser [Mar78/7]. – 1 �, 1 �; same data, 1.X.1979; leg. P. strinati.

disTRiBuTioN: Lepthyphantes aelleni is known from two potholes, the “gouffrede kaf el Bouk” and the “gouffre du Friouato”, in the region of Taza, Morocco only(denis & dresco, 1957; Bosmans, 2006). detailed information on these localities canbe found in strinati (1952).

Lepthyphantes s. lat. brevihamatus Bosmans, 1985 Figs 14-15MATeRiAl: 20 �; MoRoCCo, High Atlas Mts, “ifri el kaid” Cave, near AïtMehammed

south of Binelouidane; 1580 m a.s.l.; 5.vi.1978; leg. P. strinati [Mar 78/24]. – 11 �; “ifri elkaid” Cave; 5.vi.1978; leg. B. Hauser [Mar78/26].

disTRiBuTioN: The species has been reported from caves in the High Atlas Mtsof Morocco only (Bosmans, 1985, 2006).

Lepthyphantes s. lat. longihamatus Bosmans, 1985 Fig. 16MATeRiAl: 1 �; MoRoCCo, High Atlas Mts, “ifri el kaid” Cave, near AïtMehammed

south of Binelouidane; 1580 m a.s.l.; 5.vi.1978; leg. P. strinati [Mar 78/24].

disTRiBuTioN: The species has been reported from caves in the High Atlas Mtsof Morocco only (Bosmans, 1985, 2006).

Lepthyphantes s. lat. longipedis sp. n. Figs 17-21HoloTyPe: �; MoRoCCo, Middle Atlas Mts, Beni Mellal, Jbel ighnayene, near

ouaouizaght village, pothole Ji 11; 22.vi.1982; leg. J. delore, B. gailleton & A. gilbert.PARATyPes: 1 �, 5 �; collected together with the holotype. eTyMology: The specific epithet, an adjective, means “long-legged”, referring

to the length of legs in this species.diAgNosis: The new species is characterized by the thin and very long legs, as

well as by the small, reduced eyes in both sexes. The male can easily be recognized bythe shape of the lamella characteristica, as well as by the thick embolus with a largethumb and with a toothed base. The female is distinguished by the peculiar shape of itsposterior median plate.

desCRiPTioN: Male (holotype), partly damaged. Total length 2.30. Carapace1.25 long, 1.05 wide, pale brown, unmodified. eyes relatively small, with dark rings.Chelicerae 0.53. long, anterior margin with two large teeth and one small denticle.legs pale brown to yellow, thin and very long, most of them broken off. Fei and ii 2.75long each. Chaetotaxy unknown. Palp (Figs 17-19): Patella unmodified. Cymbium without posterodorsal outgrowth. Paracymbium with a tooth in middle part, and witha wide, short, backward-directed projection. lamella characteristica broad, its upperbranch obtuse and slightly widened distally. embolus with a large thumb and a few


strong teeth on embolic base. Abdomen 1.20 long, 0.80 wide, pale, almost white, dorsal pattern absent.

Female. Total length 3.05. Carapace 1.50 long, 1.20 wide, pale brown, unmo -dified. eyes relatively small. Chelicerae 0.90 long. legs pale brown to yellow, thin andvery long, most of them broken off. Fei 3.00, Feii 2.90 long. Abdomen 1.90 long, 1.20wide, pale, almost white, dorsal pattern absent. epigyne (Figs 20-21): Proscape broad,rounded, with a narrow base and a deep notch distally. lateral lobes reduced, stretcherlarge, oblong. Posterior median plate like a wide “v” with rounded ends.


Figs 9-13Lepthyphantes s. lat. aelleni denis in denis & dresco, 1957. (9) Right palp, retrolateral view.(10) lamella characteristica, lateral view. (11-13) epigyne, ventral, lateral and dorsal view,respectively.

ReMARks: According to the general genitalia conformation, the new species belongs to the afer species-group of Lepthyphantes s. lat. (see saaristo & Tanasevitch,1993) and seems to be most similar to L. longihamatus, but shows typical troglo bionticfeatures, i.e., small, reduced eyes, pale leg and body coloration, as well as very longlegs: leg i of the male paratype is 5 times longer than its body.

Beside that, the male of L. s. lat. longipedis sp. n. differs from L. longihamatus

by the shape of the paracymbial tooth (wide and blunt in L. s. lat. longipedis sp. n., narrow and pointed in L. longihamatus), by the shape of the lamella characteristica, as


Figs 14-16Lepthyphantes s. lat. brevihamatus Bosmans, 1985 (14-15) and L. s. lat. longihamatus Bosmans,1985 (16). (14-15) epigyne, ventral and dorsal view, respectively. (16) left palp, retrolateralview.

well as by some other details of the palp. The female of the new species differs fromthat of L. longihamatus by the shape of the proscape, which is considerably broaderthan long versus being about as long as broad in L. longihamatus. The epigene of L. s.


Figs 17-21Lepthyphantes s. lat. longipedis sp. n., � (17-19) and � paratypes (20-21). (17) left palp, retro-lateral view. (18) embolic division. (19) embolus. (20-21) epigyne, ventral and dorsal view,respectively.

lat. longipedis sp. n. also resembles to that of L. s. lat. brevihamatus, but clearly differsby the much deeper hole on the posterior edge of the proscape, as well as by the rounded ends of the v-shaped posterior median plate.

detailed information on the type locality of this species was given by gilbert(1983).


Lepthyphantes s. lat. maurusius Brignoli, 1978 TyPe MATeRiAl eXAMiNed: � holotype (MHNg); MoRoCCo, Middle Atlas Mts, “sidi

Mejbeur” Cave; 2.v.1974; leg. P. Thibaud & P. strinati.oTHeR MATeRiAl: 1 �; MoRoCCo, Middle Atlas Mts, “sidi Mejbeur” Cave, 1270 m

a.s.l.; 4.vi.1978; leg. P. strinati [Mar78/19].

disTRiBuTioN: The species is so far known only from a cave near Taza, Morocco(Brignoli, 1978). The male is still undescribed.

Lepthyphantes s. lat. taza sp. n. Figs 22-24HoloTyPe: �; MoRoCCo, Middle Atlas Mts, “ifri Tselet” Cave near Ain Teslit, Châra

region south-west of Taza, 1250 m a.s.l.; 3.vi.1978; leg. P. strinati [Mar 78/14].PARATyPes: 2 �; same data, “ifri Tselet” Cave; 3.vi.1978; leg. B. Hauser [Mar 78/15]. eTyMology: The specific epithet is a noun in opposition that refers to the

region of the type locality.

diAgNosis: The new species is characterized by the peculiar conformation ofthe epigyne, such as: the presence of a long process on each lateral wall, the absenceof the proscape, as well as the reduced, lateral lobes and stretcher.

desCRiPTioN: Female (holotype). Total length 2.50. Carapace 1.13 long, 0.88wide, pale brown, unmodified. eyes relatively small, with dark rings. Chelicerae 0.42.long, anterior margin with two teeth and a denticle; posterior margin with a very smalland poorly visible denticle. legs pale brown to yellow, relatively thin and long. leg i7.23 long (1.88+0.42+1.95+1.88+1.13), leg iv 6.09 long (1.83+0.33+1.38+1.67+0.88). Chaetotaxy. Fe i: 0-2-0-0, ii-iv: 0-0-0-0; Tii-ii: 2-2-2(3)-0, iii-iv: 2-1-1-0;Mti-iv: 1-1-1-1. Tmi 0.13. Abdomen 1.46 long, 0.83 wide, pale, almost white, dorsalpattern absent. epigyne (Figs 22-24): lateral walls very long, each one with a long apical process. Proscape and middle part of scape totally reduced. distal part of scapemassive, bucket-shaped, lateral lobes and stretcher reduced. Posterior median platedrop-shaped.

ReMARks: The general appearance of the epigyne of L. s. lat. taza sp. n. is similar to that of the cave-dwelling L. s. lat. aelleni and L. s. lat. maurusius, but theproscape in the new species is totally reduced, while in the other species the proscapeis distinct.


Megalepthyphantes brignolii sp. n. Figs 25-28HoloTyPe: �; MoRoCCo, Middle Atlas Mts, Bab bou idir, region of Taza, Ras Chiker

Cave, 1410 m a.s.l.; 30.iX.1979; leg. P. strinati & v. Aellen.PARATyPe: 1 �; from same locality, collected together with the holotype.


diAgNosis: The new species is characterized by the trifid apex of the lamellacharacteristica, as well as by the narrow, long, smoothly curved posterior median plateof the epigyne.

desCRiPTioN: Male (holotype), partly damaged and previously probably driedup; only one palp present, separated. Cephalic part of carapace with chelicerae dissected and lost, remaining part of carapace pale sandy-yellow, 1.05 wide. legs paleyellow, almost transparent, probably bleached. legs thin and relatively long, leg i10.25 long (2.80+0.50+2.60+2.80+1.55), iv 9.90 long (2.75+0.40+2.50+2.80+1.45).Chaetotaxy: Tii-ii: 2-1-1-2(3), iii-iv: 2-1-1-0; Mti-iv: 1-1-1-0. Metatarsus iv withouttrichobothrium. Tmi unknown, trichobothrium not found. Palp (Figs 25-26): Patella


Figs 22-24Lepthyphantes s. lat. taza sp. n., � paratype. (22-24) epigyne, ventral, dorsal and lateral view,respectively.


Figs 25-28Megalepthyphantes brignolii sp. n., � holotype (25-26) and � paratype (27-28). (25) Right palp,retrolateral view. (26) Part of embolic division. (27-28) epigyne, ventral and dorsal view, respectively.

dissected and lost. Tibia with a small rounded apical outgrowth. Cymbium with a posterodorsal protuberance. Paracymbium relatively large, posterior pocket trans -formed into an obtuse tooth directed upward. lamella characteristica trifurcate apically. embolus narrow, crescent-shaped, carina present. Abdomen 1.50 long, 1.10

wide, dorsally pale, with an indistinct grey median stripe flanked by grey paramedianspots connected to it with thin bands and transverse bands posteriorly.

Female. Total length 4.10. Carapace 1.50 long, 1.10 wide, brown. eyes normal.Chelicerae 0.65 long; anterior margin with two large teeth and one denticle, posteriormargin with four denticles. legs pale brown. leg i 10.15 long (2.80+0.50+2.80+2.60+1.45), iv ?, Feiv 2.50 long. Tii: 2-1-1-3(4), ii: 2-1-1-?, iii-iv: 2-1-1-0; Mti-iv:


Figs 29-32Tenuiphantes tenuis (Blackwall, 1852). (29) left palp, retrolateral view. (30) embolic division.(31-32) epigyne, ventral and dorsal view, respectively.

2-1-1-0. Tmi unknown, trichobothrium not found. Abdomen 2.65 long, 1.90 wide, dorsal pattern as in male. epigyne (Figs 27-28): Proscape strongly sclerotized, widerthan long, with a deep notch apically. lateral lobes and stretcher merged together, forming rectangular distal part of scape. Posterior median plate like a long, narrow,smoothly curved stripe.

ReMARks: The new species resembles the Algerian M. bkheitae (Bosmans &Bouragba, 1992), but males of both species differ by the shape of the palpal tibia andby the structure of the lamella characteristica. The female of the new species differs bythe tapered proscape, while in M. bkheitae the proscapus has parallel edges.

disTRiBuTioN: known from the type locality only; detailed information on itcan be found in gigon et al., 1980.

Tenuiphantes tenuis (Blackwall, 1852) Figs 29-32MATeRiAl: 1 �; MoRoCCo, Middle Atlas Mts, “ifri Tselet” Cave near Ain Teslit,

Châra region south-west of Taza, 1250 m a.s.l.; 3.vi.1978; leg. P. strinati [Mar 78/14]. – 1 �;“ifri Tselet” Cave; 3.vi.1978; leg. B. Hauser [Mar 78/15].

disTRiBuTioN: T. tenuis has an originally european-Ancient Mediterranean distribution, and was introduced to New Zealand (Millidge, 1988), to Chili andArgentina (Millidge, 1991), as well as to North America (Paquin et al., 2010). Probablythis is the first record of T. tenuis from a cave.

ACkNoWledgeMeNTsi am most grateful to all persons whose collections have been used in the

present study. i also am very much obliged to Peter J. schwendinger (MHNg) for theopportunity to work on the spider collections of the MHNg and for checking the manuscript. special thanks go to Pierre strinati (Cologny, switzerland) and BerndHauser (geneva, switzerland) for their important help in updating information on localities and collectors of the material studied, as well as to Robert Bosmans (gent,Belgium) for his helpful comments on the manuscript.

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